Estimating survival of a streamside salamander: importance of temporary emigration,capture response,and location

Estimating survival for highly secretive aquatic animals, such as stream salamanders, presents numerous challenges. Salamanders often spend a considerable time in refugia where they are difficult to capture. Few studies have calculated vital rates for stream salamanders, yet the need is substantial as they are threatened by a wide range of land-use stressors, especially urban development. In this study, we used 34 months of continuous field samples collected at an urban and undisturbed stream and robust design mark-recapture analysis to evaluate the importance of temporary emigration, capture response, and location on survival estimates of the salamander Desmognathus fuscus. We constructed a set of candidate models incorporating combinations of time- and location-varying capture and recapture probabilities, capture responses, temporary emigration, and survival estimates and ranked models using Akaike’s Information Criterion. We found strong support for month-specific capture probabilities, recapture probabilities, temporary emigration and a negative behavioral response to capture in the majority of months. We found no support for variation in capture probabilities, recapture probabilities, and temporary emigration between locations. However, we found that location strongly influenced survival estimates. Specifically, survival estimates were significantly higher at the undisturbed site than at the urban site. Our results emphasize the importance of estimating capture probabilities, recapture probabilities, capture response, and temporary emigration when evaluating survival in highly secretive aquatic animals. Failure to account for these population parameters will likely yield biased estimates of survival in freshwater animal populations.     

Capture–recapture models with heterogeneity to study survival senescence in the wild

Detecting senescence in wild populations and estimating its strength raise three challenges. First, in the presence of individual heterogeneity in survival probability, the proportion of high‐survival individuals increases with age. This increase can mask a senescence‐related decrease in survival probability when the probability is estimated at the population level. To accommodate individual heterogeneity we use a mixture model structure (discrete classes of individuals). Second, the study individuals can elude the observers in the field, and their detection rate can be heterogeneous. To account for detectability issues we use capture–mark–recapture (CMR) methodology, mixture models and data that provide information on individuals’ detectability. Last, emigration to non‐monitored sites can bias survival estimates, because it can occur at the end of the individuals’ histories and mimic earlier death. To model emigration we use Markovian transitions to and from an unobservable state. These different model structures are merged together using hidden Markov chain CMR models, or multievent models. Simulation studies illustrate that reliable evidence for survival senescence can be obtained using highly heterogeneous data from non site‐faithful individuals. We then design a tailored application for a dataset from a colony of black‐headed gull Chroicocephalus ridibundus. Survival probabilities do not appear individually variable, but evidence for survival senescence becomes significant only when accounting for other sources of heterogeneity. This result suggests that not accounting for heterogeneity leads to flawed inference and/or that emigration heterogeneity mimics survival heterogeneity and biases senescence estimates.     

Seasonal variation in terrestrial resource subsidies influences trophic niche width and overlap in two aquatic snake species: a stable isotope approach

Quantifying diet is essential for understanding the functional role of species with regard to energy processing, transfer, and storage within ecosystems. Recently, variance structure in the stable isotope composition of consumer tissues has been touted as a robust tool for quantifying trophic niche width, a task that has previously proven difficult due to bias in direct dietary analyses and difficulties in integrating diet composition over time. We used carbon and nitrogen stable isotope analyses to examine trophic niche width of two sympatric aquatic snakes, banded watersnakes Nerodia fasciata and black swamp snakes Seminatrix pygaea inhabiting an isolated wetland where seasonal migrations of amphibian prey cause dramatic shifts in resource availability. Specifically, we characterized snake and prey isotope compositions through time, space, and ontogeny and examined isotope values in relation to prey availability and snake diets assessed by gut content analysis. We determined that prey cluster into functional groups based on similarity of isotopic composition and seasonal availability. Isotope variance structure indicated that the trophic niche width of the banded watersnake was broader (more generalist) than that of the black swamp snake. Banded watersnakes also exhibited seasonal variation in isotope composition, suggesting seasonal diet shifts that track amphibian prey availability. Conversely, black swamp snakes exhibited little seasonal variation but displayed strong ontogenetic shifts in carbon and nitrogen isotope composition that closely paralleled ontogenetic shifts in their primary prey, paedomorphic mole salamanders Ambystoma talpoideum. Although niche dimensions are often treated as static, our results demonstrate that seasonal shifts in niche dimensions can lead to changes in niche overlap between sympatric species. Such short‐term fluctuations in niche overlap can influence competitive interactions and consequently the composition and dynamics of communities and ecosystems.     

Who are we sampling? Apparent survival differs between methods in a secretive species

Survival is a fundamental parameter in population dynamics with increasing importance in the management and conservation strategies of wildlife populations. Survival probability in vertebrates is usually estimated by live‐encounter data obtained by means of physical mark–capture–recapture protocols. Non‐invasive acoustic marking relying on individual‐specific features of signals has been alternatively applied as a marking technique, especially in secretive species. Nevertheless, to date no research has compared survival rate estimates obtained by acoustic and physical marking. We estimated half‐yearly and annual survival and recapture rates of a secretive and threatened passerine, the Dupont's lark Chersophilus duponti, using two separate live‐encounter data sets of males collected simultaneously by physical and acoustic marking in the same study area. The separate analysis of both methods led to different model structures, since transient individuals had to be accounted for in the acoustic marking but not in the physical marking data set. Furthermore, while reencounter probabilities did not differ between methods, survival estimates employing physical marking were lower than those obtained acoustically, especially between the postbreeding and the breeding period when the apparent survival of colour‐banded birds was twice as low as for acoustic marking. The combination of marking methods suggested the existence of different subsets of individuals differentially sampled within the population: whereas colour‐banded males seemed to represent the territorial fraction of the population, both resident and floater individuals were probably detected by acoustic marking. Using traditional mark–recapture methods exclusively could have misled our estimates of survival rates, potentially affecting prospective predictions of population dynamics. Acoustic marking has been poorly applied in mark–recapture studies, but might be a powerful complement to obtain accurate estimates of fundamental demographic parameters such as survival and dispersal.     

Species richness estimation and determinants of species detectability in butterfly monitoring programmes

Abstract 1. Species richness is the most widely used biodiversity index, but can be hard to measure. Many species remain undetected, hence raw species counts will often underestimate true species richness. In contrast, capture–recapture methods estimate true species richness and correct for imperfect and varying detectability. 2. Detectability is a crucial quantity that provides the link between a species count and true species richness. For insects, it has hardly ever been estimated, although this is required for the interpretation of species counts. 3. In the Swiss butterfly monitoring programme about 100 transect routes are surveyed seven times a year using a highly standardised protocol. In July 2003, control observers made two additional surveys on 38 transects. Data from these 38 quadrats were analysed to see whether currently available capture–recapture models can provide quadrat‐specific estimates of species richness, and to estimate species detectability in relation to transect, observer, survey, region, and abundance. 4. Species richness over the entire season cannot be estimated using current capture–recapture methods. The species pool was open, preventing use of closed population models, and detectability varied by species, preventing use of current open population models. Assuming a closed species pool during two mid‐season (July) surveys, a Jackknife capture–recapture method was used that accounts for heterogeneity to estimate mean detectability and species richness. 5. In every case, more species were present than were counted. Mean species detectability was 0.61 (SE 0.01) with significant differences between observers (range 0.37–0.83). Species‐specific detection at time t+ 1 was then modelled for those species seen at t for three mid‐season surveys. Detectability averaged 0.50 (range 0.17–0.81) for individual species and 0.65, 0.44, and 0.42 for surveys. Abundant species were detected more easily, although this relationship explained only 5% of variation in species detectability. 6. These are important, although not entirely unexpected, results for species richness estimation of short‐lived animals. Raw counts of species may be misleading species richness indicators unless many surveys are conducted. Monitoring programmes should be calibrated, i.e. the assumption of constant detectability over dimensions of interest needs to be tested. The development of capture–recapture or similar models that can cope with both open populations and heterogeneous species detectability to estimate species richness should be a research priority.     

Population Dynamics of the Critically Endangered Golden Lancehead Pitviper,Bothrops insularis: Stability or Decline?

Little is known about vital rates of snakes generally because of the difficulty in collecting data. Here we used a robust design mark-recapture model to estimate survival, behavioral effects on capture probability, temporary emigration, abundance and test the hypothesis of population decline in the golden lancehead pitviper, Bothrops insularis, an endemic and critically endangered species from southeastern Brazil. We collected data at irregular intervals over ten occasions from 2002 to 2010. Survival was slightly higher in the wet season than in the dry season. Temporal emigration was high, indicating the importance of accounting for this parameter both in the sampling design and modeling. No behavioral effects were detected on capture probability. We detected an average annual population decrease ( = 0.93, CI = 0.47–1.38) during the study period, but estimates included high uncertainty, and caution in interpretation is needed. We discuss the potential effects of the illegal removal of individuals and the implications of the vital rates obtained for the future persistence and conservation of this endemic, endangered species.     

Estimating abundance and population trends when detection is low and highly variable: A comparison of three methods for the Hermann's tortoise

Assessing population trends is a basic prerequisite to carrying out adequate conservation strategies. Selecting an appropriate method to monitor animal populations can be challenging, particularly for low-detection species such as reptiles. This study compares 3 detection-corrected abundance methods (capture–recapture, distance sampling, and N-mixture) used to assess population size of the threatened Hermann's tortoise. We used a single dataset of 432 adult tortoise observations collected at 118 sampling sites in the Plaine des Maures, southeastern France. We also used a dataset of 520 tortoise observations based on radiotelemetry data collected from 10 adult females to estimate and model the availability (g0) needed for distance sampling. We evaluated bias for N-mixture and capture–recapture, by using simulations based on different values of detection probabilities. Finally, we conducted a power analysis to estimate the ability of the 3 methods to detect changes in Hermann's tortoise abundances. The abundance estimations we obtained using distance sampling and N-mixture models were respectively 1.75 and 2.19 times less than those obtained using the capture–recapture method. Our results indicated that g0 was influenced by temperature variations and can differ for the same temperature on different days. Simulations showed that the N-mixture models provide unstable estimations for species with detection probabilities <0.5, whereas capture–recapture estimations were unbiased. Power analysis showed that none of the 3 methods were precise enough to detect slow population changes. We recommend that great care should be taken when implementing monitoring designs for species with large variation in activity rates and low detection probabilities. Although N-mixture models are easy to implement, we would not recommend using them in situations where the detection probability is very low at the risk of providing biased estimates. Among the 3 methods allowing estimation of tortoise abundances, capture–recapture should be preferred to assess population trends. © 2013 The Wildlife Society.     

Estimating Survival for Elusive Juvenile Pond-Breeding Salamanders

Juvenile vital rates have important effects on population dynamics for many species, but this demographic is often difficult to locate and track. As such, we frequently lack reliable estimates of juvenile survival, which are necessary for accurately assessing population stability and potential management approaches to conserve biodiversity. We estimated survival rates for elusive juveniles of 3 species, the ringed salamander (Ambystoma annulatum), spotted salamander (A. maculatum), and small-mouthed salamander (A. texanum), using 2 approaches. First, we conducted an 11-month (2016–2017) mark-recapture study within semi-natural enclosures and used Bayesian Cormack-Jolly-Seber models to estimate survival and recapture probabilities. Second, we inferred the expected annual juvenile survival rate given published vital rates for pre-metamorphic and adult ambystomatids assuming stable population growth. For all 3 species, juvenile survival probabilities were constant across recapture occasions, whereas recapture probability estimates were time-dependent. Further, survival and recapture probabilities among study species were similar. Post-study sampling revealed that the initial study period median estimate of annual survival probability (0.39) underestimated the number of salamanders known alive at 11 months. We therefore appended approximately 1 year of opportunistic data, which produced a median annual survival probability of 0.50, encompassing salamanders that we knew to have been alive. Calculation from literature values suggested a mean annual terrestrial juvenile ambystomatid survival probability of 0.49. Similar results among our approaches indicated that juvenile survival estimates for the study species were robust and likely comparable to rates in nature. These estimates can now be confidently applied to research, monitoring, and management efforts for the study species and ecologically similar taxa. Our findings indicated that similarly robust vital rate estimates for subsets of ecologically and phylogenetically similar species can provide reasonable surrogate demographic information that can be used to reveal key factors influencing population viability for data-deficient species. © 2020 The Wildlife Society.     

Making use of multiple surveys: Estimating breeding probability using a multievent‐robust design capture–recapture model

Increased environmental stochasticity due to climate change will intensify temporal variance in the life‐history traits, and especially breeding probabilities, of long‐lived iteroparous species. These changes may decrease individual fitness and population viability and is therefore important to monitor. In wild animal populations with imperfect individual detection, breeding probabilities are best estimated using capture–recapture methods. However, in many vertebrate species (e.g., amphibians, turtles, seabirds), nonbreeders are unobservable because they are not tied to a territory or breeding location. Although unobservable states can be used to model temporary emigration of nonbreeders, there are disadvantages to having unobservable states in capture–recapture models. The best solution to deal with unobservable life‐history states is therefore to eliminate them altogether. Here, we achieve this objective by fitting novel multievent‐robust design models which utilize information obtained from multiple surveys conducted throughout the year. We use this approach to estimate annual breeding probabilities of capital breeding female elephant seals (Mirounga leonina). Conceptually, our approach parallels a multistate version of the Barker/robust design in that it combines robust design capture data collected during discrete breeding seasons with observations made at other times of the year. A substantial advantage of our approach is that the nonbreeder state became “observable” when multiple data sources were analyzed together. This allowed us to test for the existence of state‐dependent survival (with some support found for lower survival in breeders compared to nonbreeders), and to estimate annual breeding transitions to and from the nonbreeder state with greater precision (where current breeders tended to have higher future breeding probabilities than nonbreeders). We used program E‐SURGE (2.1.2) to fit the multievent‐robust design models, with uncertainty in breeding state assignment (breeder, nonbreeder) being incorporated via a hidden Markov process. This flexible modeling approach can easily be adapted to suit sampling designs from numerous species which may be encountered during and outside of discrete breeding seasons.     

Challenges of DNA-Based Mark-Recapture Studies of American Black Bears

Abstract: We explored whether genetic sampling would be feasible to provide a region-wide population estimate for American black bears (Ursus americanus) in the southern Appalachians, USA. Specifically, we determined whether adequate capture probabilities (p > 0.20) and population estimates with a low coefficient of variation (CV < 20%) could be achieved given typical agency budget and personnel constraints. We extracted DNA from hair collected from baited barbed-wire enclosures sampled over a 10-week period on 2 study areas: a high-density black bear population in a portion of Great Smoky Mountains National Park and a lower density population on National Forest lands in North Carolina, South Carolina, and Georgia. We identified individual bears by their unique genotypes obtained from 9 microsatellite loci. We sampled 129 and 60 different bears in the National Park and National Forest study areas, respectively, and applied closed mark-recapture models to estimate population abundance. Capture probabilities and precision of the population estimates were acceptable only for sampling scenarios for which we pooled weekly sampling periods. We detected capture heterogeneity biases, probably because of inadequate spatial coverage by the hair-trapping grid. The logistical challenges of establishing and checking a sufficiently high density of hair traps make DNA-based estimates of black bears impractical for the southern Appalachian region. Alternatives are to estimate population size for smaller areas, estimate population growth rates or survival using mark-recapture methods, or use independent marking and recapturing techniques to reduce capture heterogeneity.     

Partitioning Detectability Components in Populations Subject to Within-Season Temporary Emigration Using Binomial Mixture Models

Detectability of individual animals is highly variable and nearly always < 1; imperfect detection must be accounted for to reliably estimate population sizes and trends. Hierarchical models can simultaneously estimate abundance and effective detection probability, but there are several different mechanisms that cause variation in detectability. Neglecting temporary emigration can lead to biased population estimates because availability and conditional detection probability are confounded. In this study, we extend previous hierarchical binomial mixture models to account for multiple sources of variation in detectability. The state process of the hierarchical model describes ecological mechanisms that generate spatial and temporal patterns in abundance, while the observation model accounts for the imperfect nature of counting individuals due to temporary emigration and false absences. We illustrate our model’s potential advantages, including the allowance of temporary emigration between sampling periods, with a case study of southern red-backed salamanders Plethodon serratus. We fit our model and a standard binomial mixture model to counts of terrestrial salamanders surveyed at 40 sites during 3–5 surveys each spring and fall 2010–2012. Our models generated similar parameter estimates to standard binomial mixture models. Aspect was the best predictor of salamander abundance in our case study; abundance increased as aspect became more northeasterly. Increased time-since-rainfall strongly decreased salamander surface activity (i.e. availability for sampling), while higher amounts of woody cover objects and rocks increased conditional detection probability (i.e. probability of capture, given an animal is exposed to sampling). By explicitly accounting for both components of detectability, we increased congruence between our statistical modeling and our ecological understanding of the system. We stress the importance of choosing survey locations and protocols that maximize species availability and conditional detection probability to increase population parameter estimate reliability.     

Variations in age‐ and sex‐specific survival rates help explain population trend in a discrete marine mammal population

Understanding the drivers underlying fluctuations in the size of animal populations is central to ecology, conservation biology, and wildlife management. Reliable estimates of survival probabilities are key to population viability assessments, and patterns of variation in survival can help inferring the causal factors behind detected changes in population size. We investigated whether variation in age‐ and sex‐specific survival probabilities could help explain the increasing trend in population size detected in a small, discrete population of bottlenose dolphins Tursiops truncatus off the east coast of Scotland. To estimate annual survival probabilities, we applied capture–recapture models to photoidentification data collected from 1989 to 2015. We used robust design models accounting for temporary emigration to estimate juvenile and adult survival, multistate models to estimate sex‐specific survival, and age models to estimate calf survival. We found strong support for an increase in juvenile/adult annual survival from 93.1% to 96.0% over the study period, most likely caused by a change in juvenile survival. Examination of sex‐specific variation showed weaker support for this trend being a result of increasing female survival, which was overall higher than for males and animals of unknown sex. Calf survival was lower in the first than second year; a bias in estimating third‐year survival will likely exist in similar studies. There was some support first‐born calf survival being lower than for calves born subsequently. Coastal marine mammal populations are subject to the impacts of environmental change, increasing anthropogenic disturbance and the effects of management measures. Survival estimates are essential to improve our understanding of population dynamics and help predict how future pressures may impact populations, but obtaining robust information on the life history of long‐lived species is challenging. Our study illustrates how knowledge of survival can be increased by applying a robust analytical framework to photoidentification data.     

Male-biased sex ratios in mature damselfly populations: real or artefact?

1. There is ongoing controversy about whether biased sex ratios in diploid insect populations are real or an artefact caused by different behaviours and/or different catchability of the sexes. This was tested by monitoring two field and three semi-natural populations of the damselfly Lestes sponsa. 2. Capture–mark–recapture data showed that population size estimates were about twice as large for males as for females at both field sites. Independent estimates of the sex ratios based on total numbers of males and females captured supported the male bias. 3. Males had higher recapture probabilities than females due to longer times between successive visits in females. Because the same pattern was found in the semi-natural populations, the longer intervals in females are no artefact due to their lower detectability. 4. Theoretical models show that the strong temporary emigration of females tends, if anything, to overestimate female population sizes and that the heterogeneity of recapture probabilities observed in males tends to underestimate male population sizes. Hence, behavioural differences between the sexes do not cause an artificially male-biased sex ratio. 5. Spatial data show that operational sex ratios are male biased at the pond but become female biased in the plots further away from the shoreline; however because of the decrease in densities away from the shoreline, this does not result in a global even sex ratio. 6. Spatial data, temporary emigration patterns, and independent estimates suggest strongly that the male-biased sex ratios in mature damselfly populations are real.     

Seasonal abundance and adult survival of bottlenose dolphins (Tursiops truncatus) in a community that cooperatively forages with fishermen in southern Brazil

A subgroup of a population of Tursiops truncatus in southern Brazil is known for a cooperative behavior with artisanal fishermen whereby the dolphins shoal fish towards net‐casting fishermen. Combining photo‐identification data collected between September 2007 and 2009 with mark‐recapture and Pollock's robust design models, we assessed abundance within seasons and survival and temporary emigration rates of dolphins between seasons. We also reanalyzed a previous data set collected during 1989–1991, and Cormack‐Jolly‐Seber models were applied to estimate survival rates for each of the study periods. The abundance of marked “cooperative” dolphins varied between seasons from 18 (CI: 17–24) to 21 (CI: 20–24). The total abundance varied from 59 in the winter of 2008 (CI: 49–72) to 50 in the autumn of 2009 (CI: 40–62). The annual adult survival was estimated to be 0.917 (CI: 0.876–0.961), close to that estimated from data collected in the 1990s (0.941; CI: 0.888–0.998). The emigration probability was low (0.031; CI: 0.011–0.084) and different capture probabilities between the “cooperative” and “noncooperative” dolphins indicated a degree of behavioral segregation. The precision of our estimates is likely to provide sufficient power to detect population change, but we recommend a precautionary management approach to protect this vulnerable dolphin community and its unique cooperative feeding tradition.     

Movement patterns and study area boundaries: influences on survival estimation in capture–mark–recapture studies

The inability to account for the availability of individuals in the study area during capture–mark–recapture (CMR) studies and the resultant confounding of parameter estimates can make correct interpretation of CMR model parameter estimates difficult. Although important advances based on the Cormack–Jolly–Seber (CJS) model have resulted in estimators of true survival that work by unconfounding either death or recapture probability from availability for capture in the study area, these methods rely on the researcher's ability to select a method that is correctly matched to emigration patterns in the population. If incorrect assumptions regarding site fidelity (non‐movement) are made, it may be difficult or impossible as well as costly to change the study design once the incorrect assumption is discovered. Subtleties in characteristics of movement (e.g. life history‐dependent emigration, nomads vs territory holders) can lead to mixtures in the probability of being available for capture among members of the same population. The result of these mixtures may be only a partial unconfounding of emigration from other CMR model parameters. Biologically‐based differences in individual movement can combine with constraints on study design to further complicate the problem. Because of the intricacies of movement and its interaction with other parameters in CMR models, quantification of and solutions to these problems are needed. Based on our work with stream‐dwelling populations of Atlantic salmon Salmo salar, we used a simulation approach to evaluate existing CMR models under various mixtures of movement probabilities. The Barker joint data model provided unbiased estimates of true survival under all conditions tested. The CJS and robust design models provided similarly unbiased estimates of true survival but only when emigration information could be incorporated directly into individual encounter histories. For the robust design model, Markovian emigration (future availability for capture depends on an individual's current location) was a difficult emigration pattern to detect unless survival and especially recapture probability were high. Additionally, when local movement was high relative to study area boundaries and movement became more diffuse (e.g. a random walk), local movement and permanent emigration were difficult to distinguish and had consequences for correctly interpreting the survival parameter being estimated (apparent survival vs true survival).     

Effect of Tooth Removal on Recaptures of Raccoons

Abstract: Researchers have extensively used mark—recapture techniques to obtain information on demographic parameters of wildlife populations. However, researchers have recognized that a number of factors can influence capture probabilities of wildlife species, which in turn can bias mark—recapture estimates of demographic parameters. Tooth extraction, which is a commonly used technique in studies of mesopredator species to obtain precise age estimates and to monitor the use of vaccine baits, is an aspect of animal handling that clearly might affect the recapture probability of individuals. However, the effect that tooth removal has on the individual recapture probabilities of wildlife species is unknown. During 2005, we trapped and marked 91 raccoons (Procyon lotor) in northern Indiana, USA, as part of a mark—recapture study designed specifically to determine if tooth extractions have an effect on recapture probabilities of individuals. We performed tooth extractions on 50% of the raccoons at the time of capture, and we attempted to balance tooth extractions with respect to sex and age of raccoons. We used logistic regression to model the effects of sex, age, and tooth removal on recapture probabilities, and we used Mann—Whitney U-tests to examine the effect of tooth removal on the number of times we recaptured individuals. The probability of recapture differed between sexes but did not differ as a function of tooth removal or among age classes. In addition, we failed to detect any difference in the mean number of times that we recaptured raccoons between the tooth removed and non—tooth-removed groups. Our results suggest that managers can use tooth extractions as an effective management tool without biasing population estimates or compromising other management objectives.     

Effects of body size,diet, and transience on the demography of the arboreal boid snake Corallus grenadensis on Carriacou (Grenada Grenadines,West Indies)

Remarkably little is known about the demography of snakes in the family Boidae. This lack of information may be attributed, in part, to low population densities on the Neotropical mainland, rendering capture-recapture methods impractical for many species. Conversely, islands support fewer species but snake densities can be much higher. Corallus grenadensis is an arboreal boid endemic to the Grenada Bank and, relative to mainland boids, can be amazingly abundant. As young, its diet is comprised largely of native Anolis lizards, a ubiquitous and abundant food source; it then undergoes an ontogenetic shift in diet to a less abundant resource, rodents. From 2015 to 2019, we marked 254 C. grenadensis and used capture–recapture models to estimate abundance, capture probabilities, survival, and the proportion of transients. We hypothesized that the transient effect would increase with body size (snout–vent length [SVL]), prompted by their ontogenetic shift in diet. Capture probabilities increased with sampling effort and decreased with increasing SVL. Abundance ranged from 96 to 141 individuals and annual resident survival was 0.71, 95% confidence interval (CI) = 0.54–0.83. The proportion of transients increased with increasing SVL, with the estimate being distinguishable from zero starting at ~810 mm SVL, coinciding with the size at which their dietary shift from ectothermic to endothermic prey begins. Ontogenetic dietary shifts are widespread in snakes and occur in at least 11 of 17 species of West Indian boids. Thus, the prominence of transients in our study may be indicative of its demographic and ecological importance among other snake species.     

Estimating Detection Probabilities of Waterfowl Broods From Ground-Based Surveys

ABSTRACT Brood:pair ratios could provide an economical method for assessing spatial or temporal variation in waterfowl productivity, but such estimators are severely biased by incomplete detection of broods. We conducted 3 sequential counts of 1,357 waterfowl broods in northeastern North Dakota, USA, and used closed-population mark-recapture models to estimate total brood abundance while controlling for variation in detection probabilities (p). Blue-winged teal (Anas discors) broods had the lowest average detection probability (p = 0.305), whereas diving-duck broods had the highest average detectability (p = 0.571). Detection was generally highest in morning or evening, but temporal patterns varied among species and there was no survey window that maximized detection probabilities for all species. Detection probabilities averaged 0.108 (SD = 0.056) higher for an experienced observer versus an inexperienced observer. Detection probabilities were 0.044 higher for roadside versus walk-up surveys and increased with increasing brood size, total brood abundance, survey date, wind speed, temperature, cloud cover, and amount of time spent surveying each wetland. Detection probabilities declined with increasing wetland size and amount of tall peripheral vegetation. Our mark-recapture results indicated that a traditional unreplicated brood survey would have missed 67.5% of estimated broods, summed over all species. Use of closed-population mark-recapture techniques provided an effective method for reducing this bias and identifying and quantifying factors that reduce detection probabilities of waterfowl broods. We recommend that future brood surveys incorporate 2 or 3 temporally segregated replicate counts to allow for formal estimation of detection probabilities.     

Loglinear models for the robust design in mark-recapture experiments

The robust design is a method for implementing a mark-recapture experiment featuring a nested sampling structure. The first level consists of primary sampling sessions; the population experiences mortality and immigration between primary sessions so that open population models apply at this level. The second level of sampling has a short mark-recapture study within each primary session. Closed population models are used at this stage to estimate the animal abundance at each primary session. This article suggests a loglinear technique to fit the robust design. Loglinear models for the analysis of mark-recapture data from closed and open populations are first reviewed. These two types of models are then combined to analyze the data from a robust design. The proposed loglinear approach to the robust design allows incorporating parameters for a heterogeneity in the capture probabilities of the units within each primary session. Temporary emigration out of the study area can also be accounted for in the loglinear framework. The analysis is relatively simple; it relies on a large Poisson regression with the vector of frequencies of the capture histories as dependent variable. An example concerned with the estimation of abundance and survival of the red-back vole in an area of southeastern Québec is presented.     

Augmenting superpopulation capture-recapture models with population assignment data

Summary Ecologists applying capture–recapture models to animal populations sometimes have access to additional information about individuals' populations of origin (e.g., information about genetics, stable isotopes, etc.). Tests that assign an individual's genotype to its most likely source population are increasingly used. Here we show how to augment a superpopulation capture–recapture model with such information. We consider a single superpopulation model without age structure, and split each entry probability into separate components due to births in situ and immigration. We show that it is possible to estimate these two probabilities separately. We first consider the case of perfect information about population of origin, where we can distinguish individuals born in situ from immigrants with certainty. Then we consider the more realistic case of imperfect information, where we use genetic or other information to assign probabilities to each individual's origin as in situ or outside the population. We use a resampling approach to impute the true population of origin from imperfect assignment information. The integration of data on population of origin with capture–recapture data allows us to determine the contributions of immigration and in situ reproduction to the growth of the population, an issue of importance to ecologists. We illustrate our new models with capture–recapture and genetic assignment data from a population of banner‐tailed kangaroo rats Dipodomys spectabilis in Arizona.